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CLINTON NATHAN WOOLSEY November 30. ~ 904-fanuary ~ 4, ~ 993 BY RICHARD F. THOMPSON ONE OF THE GREAT achievements of neuroscience in this century has been characterization of the organization of sensory en c! motor representations in the cerebral cor- tex. Rough facts of cortical localization were known from the nineteenth century en c! earlier, baser! on studies from brain-ciamagec! humans en c! animals en c! by application of electrical stimulation. The most important contemporary scientist to pioneer the fine-grainec! analysis of sensory en c! motor representations in the cortex was Clinton N. Woolsey. In the course of his remarkable career he macle many im- portant en c! funciamental discoveries. Clinton N. Woolsey was born on November 30, 1904, in Brooklyn, New York, the son of Joseph Wooc~hull en c! Mathilcia Louise Aicho~z Woolsey. He left Brooklyn at the age of nine months ("before cleveloping a Brooklyn accent". He spent his youth in Orange County, New York, en c! attenclec! a one- room country school from gracles ~ through 6. He clescribec! it as an interesting experience, because he "conic! listen to the lessons given to all the pupils." He attended grades 7 through ~ O in Montgomery, New York, ant! mover! to Schenevus, New York, for his thirc! year of high school. There he was awarclec! the H. Bernarc! GoIc! Mecial as the best student of the year. 361

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362 BIOGRAPHICAL MEMOIRS In the audience cluring his aware! ceremony was Dr. Walter AlIan Cowell of Olean, New York, who suggestec! that young Woolsey move to Olean for his senior year, which he clicI. Cowell was a physician interested both in the practice of medicine en c! in research en c! was studying the effects on diabetes of insulin, which hac! just been cliscoverecI. As a result of his association with Cowell, Woolsey became greatly interested in medicine en c! in research. He gracluatec! near the top of his class en c! spent another year at Olean High School taking extra work in Latin, French, en c! other subjects. In 1924 Woolsey enterer! Union College in Schenectady, New York, where Cowell hac! gracluatecI. Woolsey continues! his study of Latin en c! French en c! took two years of Greek en c! the courses neeclec! to enter meclical school. Among his most impressionable experiences at Union College was a remarkable psychology professor, Johnny March, who "cle- scribed the experiments of PavIov and Sherrington so viv- idly that one felt in the presence of these investigators and their experimental animals." As a result of this, Woolsey consiclerec! going to Columbia University for training in psychology. Instead, he decided to go to medical school, and was accepted by the Johns Hopkins University School of Medicine in ~ 928. During his first year at Hopkins, Woolsey took courses in histology en c! neuroanatomy from Dr. Marion Hines. At that time Woolsey intenclec! to become a brain surgeon, so Hines sent him to work with Dr. Sarah Tower, who was an accom- plished animal surgeon. Following a special course on To- calization of function taught jointly by Hines en c! Tower, Hines invites! Woolsey to work with her on the clog brain, which led to his first publication: "On the Postural Rela- tions of the Frontal and Motor Cortex of the Dog" (1933~. Before finishing his fourth year of medical studies, Woolsey clevelopec! pulmonary tuberculosis (a not uncommon con

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CLINTON NATHAN WOOLSEY 363 dition for medical students of the day) and had to leave school for six months to recuperate in a sanitarium. He was advised that his goal of an internship in surgery was too physically demanding and might reactivate his pulmonary lesion. Dr. Philip Bard had just been recruited to Hopkins from Harvard, and he invited Woolsey to work in his lab. Woolsey soon realized that his future lay in physiology and not brain surgery. (In later years, those of us who worked with him through all-night experiments and 14-hour brain surgeries on monkeys were amazed at his enormous energy and robust health. ) During his time at Hopkins, Woolsey married Harriet E. Runion (in ~ 942) . They had three children: Thomas Allen Woolsey, M.D., now a leading neuroscientist, John David Woolsey, M.F.A., an artist and medical illustrator, and Ed- ward Alexander Woolsey, Ph.D., a zoologist and teacher. . . . Woolsey remained at Hopkins in physiology until 194S, when he accepted his appointment as Charles Sumner Slichter professor of neurophysiology at the University of Wisconsin medical and graduate schools in Madison. He remained at Wisconsin for the rest of his career and life. I had the great good fortune to work in Clinton Woolsey's laboratory for four years from 1955 through 1959. I com- pleted by Ph.D. thesis in 1955-56 in his laboratory (my ma- jor professor, W. l. Brogden, in the psychology department did not have facilities for my work, and Woolsey kindly al- lowed us to use his laboratory). I then spent three years in Woolsey's laboratory as an NIH postdoctoral fellow. It was a most exciting environment. Much of the work in the labo- ratory at the time focused on the organization of the motor cortex in a series of primates (including chimpanzees) us- ing electrical stimulation and on the organization of sen- sory (and polysensory) cortical areas using surface-evoked potentials. P. W. Davies visited the lab during that time and

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364 BIOGRAPHICAL MEMOIRS clescribec! the new extracellular microelectrocle technique he en c! {erzy Rose hac! clevelopecI. {erzy Rose visitor! one summer, en c! using this technique we completec! the first single-unit recording study of the tonotopic organization of the auditory cortex (in cat) (1960~. During my time (and, of course, earlier and later, as well) there were extraorclinariTy taTentec! scientists in the labora- tory. Konrac! Akert proviclec! solic! expertise in neuroanatomy (e.g., 1961), Joseph Hint! was expert in the auditory system en c! all matters acoustic (1960), en c! W. I. Welker en c! Rob ert Benjamin were young scientists at the height of their productivity (e.g., 1957~. There were many others as well (e.g., 1957~. Woolsey was a very tolerant laboratory chief. If the work we clic! was to some degree relevant to cortical organization en c! functions en c! was carefully clone, we were free to follow our own interests. Personally, Woolsey was a gentle man I never saw him lose his temper. He was an icleal role moclel in that he was totally focuses! on the work (anc! his family), was objective, en c! never engages! in act hominem. However, if you took a particular position on cortical organization, you hac! better be preparer! to clefenc! it. He had very high standards and expected the same of everyone. Morale in Woolsey's laboratory was extremely high. Woolsey was a superb but infrequent lecturer, often teach- ing by demonstration. At that time textbooks states! that complete removal of the neocortex in monkeys causer! vir- tual paralysis. In the meclical student physiology course Woolsey once clemonstratec! a fully clecorticate rhesus mon- key, which he hell! on a stick chain while the monkey chaser! him arounc! the lectern trying to bite him. This fincling J ~ was, of course, much more than simply a demonstration. Travis en c! Woolsey (1956) shower! that after bilateral re moval of all neocortex in stages, monkeys conic! show con siclerable recovery of motor function en c! become capable

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CLINTON NATHAN WOOLSEY 365 of locomotion if given adequate postoperative physical therapy. Recovery of function following brain injury was of creep interest to Woolsey. I assistec! Woolsey in preparation of two of the clecorticate macaques (we clic! them in two stages) en c! in their postoperative care. Woolsey hac! clevel- opec! a methoc! of subpial surgical aspiration of cortex that macle it possible to remove TocaTizec! regions without ciam- age to adjacent regions, or of an entire hemisphere of cor- tex with minimal bleecling. He was a superb experimental neurosurgeon. Woolsey became the Charles Sumner Slichter professor emeritus at the University of Wisconsin in 1975, but he by no means retiree! from his work. He publisher! a lancimark paper on localization in somatic sensory en c! motor areas of the human cerebral cortex in 1979, en c! until shortly before his cleath he was hare! at work bringing to comple- tion his extensive ciata on cortical localization in the chim- panzee. I participates! in these studies, en c! he caller! me, I believe in 1990, to check on some cletaiTs. In the course of his long en c! productive career, Clinton Woolsey receiver! many honors en c! awards, inclucling Phi Beta Kappa (1928), the Franklin P. Mall Awarc! in anatomy, Johns Hopkins University ( 1933), National Academy of Sci- ences membership (1960), the Mecial of Faculty of Mecli- cine, Free University, Brussels, Belgium ~ ~ 968), charter membership in the Johns Hopkins Society of Scholars (1968), an Sc.D. (honoris cause) from Union College (1968), hon- orary membership in the Academy of Neurosurgery ~ ~ 973), honorary membership in the American Neurological Asso- ciation (1975), en c! the Ralph W. Gerarc! Awarc! from the Society for Neuroscience, with l. F. Rose (1982~. He server! on numerous NIH committees en c! was cleeply involves! in international scientific activities.

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366 BIOGRAPHICAL MEMOIRS PROFESSIONAL HISTORY Wacle Marshall hac! joiner! Philip Barcl's department at Johns Hopkins in the 1930s en c! hac! worker! with the cath- ocle ray oscilloscope in Ralph Gerarcl's laboratory. He en c! Albert Grass at Harvarc! built such equipment for Barcl's laboratory, en c! Marshall, Woolsey, en c! B arc! undertook the first cletailec! mapping of the somatic sensory area of the cerebral cortex of the cat en c! monkey, using the new evokoc! potential technique (1937, 1941, 1942~. This formed the basis of much of Woolsey's future work. He mapped the cortical sensory areas (anc! motor areas) in many mamma- lian species in great cletail. Woolsey hac! a creep en c! abicling interest in the comparative clevelopment of functional ar- eas of the neocortex, an interest he conveyor! to his student en c! colleague W. I. Welker, who has continues! this impor- tant tradition to the present. These studies were clone with great care and attention to detail and led to a general for- mulation in the late 1940s of the receptotopic organization of cortical receiving areas, the general plan of which is largely unchallengec! to this clay. The power of this comparative approach is clear in the following passage: Of particular significance in the evolution of these [somatic sensory mo- tor] fields is the central position of the hand areas of SI and MI. In the primates the hand achieves a high degree of corticalization in the precen- tral and the postcentral fields. Because of the central location of the hand areas, the simple basic pattern of organization seen in the rodent, where the parts are represented in relation to one another much as they exist in the actual animal, apparently becomes distorted in evolution as cortical representation for the hand increases, with the result that in chimpanzee and in man the sensory and the motor face areas lose continuity with the centers for occiput and neck, which remain associated with the trunk rep- resentations. In macaque this separation of face from occiput has taken place in the postcentral gyrus, but in the precentral field the motor pattern still hangs together as it does in lower forms. Evidence for a transitional

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CLINTON NATHAN WOOLSEY 367 status in the postcentral area in the smooth-brained marmoset has been reported and illustrated elsewhere. That this separation of cortical centers for face and occiput is not the result of an en bloc reversal of the projec- tions of the cervical segments upon the cortex as was once suggested (1942), but rather is due to expansion of the hand area and disruption thereby of the cortical pattern, is supported by the finding that the trigeminal nerve projects not only to the lower classical face area but also to the "upper" head area, where not only the occiput but other parts of the head and face are represented ( 1958, pp. 65-66~ . In his initial studies Woolsey focuses! on the somatic sen- sory cortex, but he quickly extenclec! the work to auditory en c! visual areas. It was known that regions of the cochlea responclec! selectively to different tone frequencies, but little was known about the auditory cortex. Woolsey en c! Walz! (1942) completec! a technical tour cle force by selectively stimulating TocaTizec! regions of auditory nerve fibers in the cochlea en c! mapping the patterns of evokoc! responses on the auditory cortex of the cat en c! monkey. This was the first clear demonstration of tonotopic (actually cochicotopic) organization of the auditory cortex. They follower! this by examining effects of cochIear lesions on click-evokoc! re- sponses in the auditory cortex (1946~. Early in the 1940s Woolsey cliscoverec! the existence of a seconc! somatic sensory receiving area in the cortex of the cat, clog, en c! monkey ~ ~ 943) en c! subsequently cliscoverec! secondary auditory en c! visual areas. Both E. D. Adrian en c! Woolsey are creclitec! with inclepenclent discovery of the ex- istence of this seconc! somatic sensory area. Actually, it ap- pears that Woolsey was first. The following is a quote from a letter written to me by Clinton's son Thomas: I believe Dad and his colleagues in Baltimore discovered a second somatic area independently and about the same time as Adrian. My father rarely expressed disappointments in others. However, I think this is one case where he was both very disappointed and surprised. Evidently, early during the Second World War, Adrian was in the United States and visited the

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368 BIOGRAPHICAL MEMOIRS laboratory in Baltimore, Maryland. As Father described it, he [Clinton Woolsey] spent a long time very carefully explaining his discovery of a second so- matic area. Father said that Adrian nodded and made comments regarding the data that he was being shown, but said nothing about his own work, which Father was greatly surprised to see published several months later. I think Dad felt betrayed in his confidence. In any case, a review of the data suggests that Dad provided the first convincing evidence of an orderly sequence in a second full representation. Adrian's note had only a few points that were outside the region of what was already known to be the somatic area (SI)" (Thomas Woolsey, personal communication, Nov. 22, 1994~. Woolsey en c! associates carefully mapper! the primary vi- sual area of the cortex, clemonstratec! the cletailec! retinotopic organization, en c! mapper! a seconc! visual area ~ ~ 946, ~ 950) . In yet another series of pioneering studies, Woolsey en c! associates mapper! the somatic sensory projections to the cerebelIar cortex (1945) en c! the organization of projec- tions from the cerebral cortex to the cerebellar cortex (1952~. In still yet another series of pioneering studies, Woolsey joiner! forces with {erzy Rose to complete a cletailec! lesion retrograde degeneration mapping of the projections from the auditory region of the thalamus (meclial geniculate bocly) to the auditory cortex in light of the physiological organi- zation of the auditory cortex Woolsey en c! Walz! hac! cle- finec! earlier (1949~. Rose en c! Woolsey completec! similar studies on the projections of the meclioclorsal nucleus to the orbitofrontal cortex (1947) en c! on the relations be- tween the anterior thalamic nuclei en c! the limbic cortex (1948~. As notes! by Clinton's son Thomas in the presenta- tion statement for the Ralph W. Gerard Award to Woolsey en c! Rose in 1982, these studies clemonstratec! that (~) there was a direct correspondence between cortical cytoarchitec- tonic fielcis en c! functionally clefinec! regions of the cortex (this structure-function concept was under attack at the time), (2) each functional en c! cytoarchitectonic region of cortex

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CLINTON NATHAN WOOLSEY 369 receiver! a distinctive input from a specific thalamic nucleus (the concept of thalamotelencephalic clepenclencies), en c! (3) these connections either conic! be restrictec! or be clis- tributec! more wiclely to several functional en c! cytoarchitec- tonic areas (the concepts of essential en c! sustaining projec- tions) . In an ongoing series of exquisitely cletailec! studies, Woolsey en c! associates mapper! the primary en c! supplementary motor areas of the cerebral cortex, using electrical stimulation in a wicle range of primates en c! other mammals (1952, 1957, 1958) en c! comparer! sensory en c! motor maps in both pre- anc! postcentral cortical areas: It has now been firmly established that the afferent areas are not strictly afferent nor are the motor areas entirely motor. The afferent areas (SI and SII; postcentral and "second" sensory) have well-organized motor outflows which are still functional months after complete removal of the motor areas of the frontal lobe, while at the same time it appears that afferent connections to the frontal motor areas exist independently of the parietal afferent paths (Figure 1~. Thus, the concept that the rolandic region is indeed a sensorimotor system, as held by pre-Sheringtonian workers, is reaffirmed, but with the considerable difference that the region is not an undifferentiated entity but one compounded of a number of distinguish- able, individually complete, though interrelated, sensory-motor and motor- sensory representations. These facts appear to us to have important conse- quences for studies of the role of the cortex in neurological and behavioral functions, studies which will require the close cooperation of anatomist, physiologist, and behaviorist, or the mastery of multiple techniques by single individuals ( 1958, p. 64~ . It is perhaps fitting to close this review of Clinton Woolsey's professional history with an example of his work. Figure ~ is reproclucec! from Woolsey ~ ~ 958) . It shows the cletailec! maps of a portion of the postcentral gyrus of the Macaca mulatto, comparing the representation of the belly surface on the left, obtainer! from evokes! potential maps, to the representation of movements from the same cortical tissue,

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370 BIOGRAPHICAL MEMOIRS ~N t;~t,~v\'! { ~ tofu /~' / : \ ~.~\ /~p / /6 ~ / / /6~7~/ J ~/ /~1~ ,~,~~1 Uj - ~' .~1~,~8~) ~' / / POSTCENTRAL ~ - ' - POSTCENTRAL TACTILE ~MOTOR FIGURE 1: Comparison of postcentral tactile localization pattern with the postcen- tral motor localization pattern of Macaca mulatto (1958). elicitec! by electrical stimulation, on the right. The figu- rines (a style of data representation Woolsey invented and perfected) in both cases indicate 2-mm steps along the cor- text The dark region in each figurine on the left is the region of skin surface that yielcis the maximum amplitucle evokes! potential at that cortical locus when the skin is lightly tapped. The dark regions on the figurines on the right are the cortical loci where least-intensity electrical stimulation yielcis the minimal movement shown. Note the exquisite cletail. In the animal on the right (motor map), the cortical motor area (precentral gyrus) had been removed bilater- ally, hence there is a well-organized postcentral motor sys- tem that can function independently of the frontal motor paths (see quotation just above). Woolsey also notes that

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CLINTON NATHAN WOOLSEY 371 these two maps are clerivec! from different animals, yet they show remarkable similarities in their patterns of somato- topical organization. It wouic! be very remiss of me to conclucle this biography of Clinton Woolsey without special mention of his wife Harriet. She was totally supportive of his work en c! career en c! acceptec! with profounc! goof! nature his increclible work scheclule, he wouIc! often work all clay en c! night en c! some- times longer. Clinton was truly most fortunate. We who worker! with him remember with great fanciness the eve- nings at their home. Every summer, when the first sweet corn was ripe in southern Wisconsin, Clinton en c! Harriet hell! a corn roast for the laboratory at a local park. Perhaps it was the influence of Harriet en c! Clinton, but somehow corn has never taster! quite as goof! since. ~ AC~OWLEDGE MOST gratefully the following documents that pro- vided information, particularly on the early phases of Clinton Woolsey's life: the autobiographical document dated April 10,1989, that Clinton Woolsey wrote to the National Academy of Sciences; the original nomination (circa 1959) to the National Academy of Sciences; the Ralph W. Gerard Award presentation statement dated November 1, 1982, that Clinton's son Thomas A. Woolsey wrote for the Society for Neuroscience; and a letter dated November 22, 1994, that Tho- mas A. Woolsey wrote to me. Finally, I have innumerable personal experiences from the time I worked in Clinton Woolsey's laboratory from 1955 to 1959, the high point of my professional career.

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372 BIOGRAPHICAL MEMOIRS SELECTED BIBLIOGRAPHY 1937 With W. H. Marshall and P. Bard. Cortical representation of tactile sensibility as indicated by cortical potentials. Science 85:388-90. 1941 With W. H. Marshall and P. Bard. Observations on cortical somatic sensory mechanisms of cat and monkey. 7. Neurophysiol. 4:1-43. 1942 With W. H. Marshall and P. Bard. Representation of cutaneous tac- tile sensibility in the cerebral cortex of the monkey as indicated by evoked potentials. Bull. fohns Hopkins Hosp. 70: 399-441. With E. M. Walzl. Topical projection of nerve fibers from local regions of the cochlea to the cerebral cortex of the cat. Bull. fohns Hopkins Hosp. 71:315-44. 1943 "Second" somatic receiving areas in the cerebral cortex of cat, dog, and monkey. Fed. Proc. 2:55-56. 1945 With J. L. Hampson and C. R. Harrison. Somatotopic localization in the anterior lobe and lobulus simplex of the cerebellum of the cat and dog. Fed. Proc. 4:31. i946 With S. A. Talbot and J. M. Thompson. Visual areas I and II of the cerebral cortex of the rabbit. Fed. Proc. 5:103. With E. M. Walzl. Effects of cochlear lesions on click responses in the auditory cortex of the cat. Bull. fohns Hopkins Hosp. 79:309- 19. 1947 With J. E. Rose. The orbitofrontal cortex and its connections with the mediodorsal nucleus in rabbit, sheep, and cat. Res. Publ. Assoc. Nero. Ment. Dis. 27:210-32.

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CLINTON NATHAN WOOLSEY 1948 373 With J. E. Rose. Structure and relations of limbic cortex and ante- rior thalamic nuclei in rabbit and cat. 7. Comp. Neurol. 89:279- 348. 1949 With J. E. Rose. The relations of thalamic connections, cellular structure and evocable electrical activity in the auditory region of the cat. 7. Comp. Neurol. 91:441-46. 1950 With J. M. Thompson and S. A. Talbot. Visual areas I and II of the cerebral cortex of the rabbit. 7. Neurophysiol. 13: 277-88. 1952 Patterns of localization in sensory and motor areas of the cerebral cortex. In The Biology of Mental Health and Disease, pp. 192-206. New York: P. Hoeber. With P. H. Settlage, D. R. Meyer, W. Sencer, T. P. Hamuy, and A. M. Travis. Patterns of localization in precentral and "supplemen- tary" motor areas and their relation to the concept of the premotor area. Res. Publ. Assoc. Nerv. Ment. Dis. 30:238-64. With J. L. Hampson and C. R. Harrison. Cerebro-cerebellar projec- tions and somatotopic localization of motor function in the cer- ebellum. Res. Publ. Assoc. Nerv. Ment. Dis. 30:299-316. 1956 With A. M. Travis. Motor performance of monkeys after bilateral partial and total cerebral decortications. Am. f. Phys. Med. 35:273- 310. 1957 With W. S. Coxe, J. F. Hirsch, R. M. Benjamin. W. I. Welker, and R. F. Thompson. Precentral and supplementary motor areas in Ateles. Physiologist 1:1 9. With W. I. Welker, R. M. Benjamin, and R. C. Miles. Motor effects of cortical stimulation in squirrel monkey. J. Neurophysiol. 20:347- 64.

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374 BIOGRAPHICAL MEMOIRS 1958 With H. F. Harlow. Eds. Biological and Biochemical Bases of Behavior. Madison: University of Wisconsin Press. With J. E. Rose. Cortical connections and functional organization of the thalamic auditory system of the cat. In Biological and Bio- chemical Bases of Behavior, eds. H. F. Harlow and C. N. Woolsey, pp. 127-50. Madison: University of Wisconsin Press. Organization of somatic sensory and motor areas of the cerebral cortex. In Biological and Biochemical Bases of Behavior, eds. H. F. Harlow and C. N. Woolsey, pp. 63-81. Madison: University of Wisconsin Press. 1960 With J. E. Hind, J. E. Rose, P. W. Davies, R. M. Benjamin, W. I. Welker, and R. F. Thompson. Unit activity in the auditory cortex. In Neural Mechanisms of the Auditory and Vestibular Systems, eds. G. L. Rasmussen and W. F. Windle, pp. 201-10. Springfield, Ill.: Charles C. Thomas. Organization of the cortical auditory system: A review and a synthe- sis. In Neural Mechanisms of the Auditory and Vestibular Systems, eds. G. L. Rasmussen and W. F. Windle, pp. 165-80. Springfield, Ill.: Charles C. Thomas. 1961 With K. Akert, R. A. Grusen, and D. R. Meyer. Kluver-Bucy syn- drome in monkeys with neocortical ablations of temporal lobe. Brain 84:480-98. 1979 With T. C. Erickson and W. E. Gilson. Localization in somatic sen- sory and motor areas of human cerebral cortex as determined by direct recording of evoked potentials and electrical stimulation. 7. Neurosurg. 51:476-506.